Traps of carnivorous pitcher plants as a habitat: composition of the fluid, biodiversity and mutualistic activities

Abstract

Background: Carnivorous pitcher plants (CPPs) use cone-shaped leaves to trap animals for nutrient supply but are not able to kill all intruders of their traps. Numerous species, ranging from bacteria to vertrebrates, survive and propagate in the otherwise deadly traps. This paper reviews the literature on phytotelmata of CPPs.

Pitcher: Fluid as a Habitat The volumes of pitchers range from 0·2 mL to 1·5 L. In Nepenthes and Cephalotus, the fluid is secreted by the trap; the other genera collect rain water. The fluid is usually acidic, rich in O(2) and contains digestive enzymes. In some taxa, toxins or detergents are found, or the fluid is extremely viscous. In Heliamphora or Sarracenia, the fluid differs little from pure water.

Inquiline: Diversity Pitcher inquilines comprise bacteria, protozoa, algae, fungi, rotifers, crustaceans, arachnids, insects and amphibia. The dominant groups are protists and Dipteran larvae. The various species of CPPs host different sets of inquilines. Sarracenia purpurea hosts up to 165 species of inquilines, followed by Nepenthes ampullaria with 59 species, compared with only three species from Brocchinia reducta. Reasons for these differences include size, the life span of the pitcher as well as its fluid. MUTUALISTIC: Activities Inquilines closely interact with their host. Some live as parasites, but the vast majority are mutualists. Beneficial activities include secretion of enzymes, feeding on the plant's prey and successive excretion of inorganic nutrients, mechanical break up of the prey, removal of excessive prey and assimilation of atmospheric N(2).

Conclusions: There is strong evidence that CPPs influence their phytotelm. Two strategies can be distinguished: (1) Nepenthes and Cephalotus produce acidic, toxic or digestive fluids and host a limited diversity of inquilines. (2) Genera without efficient enzymes such as Sarracenia or Heliamphora host diverse organisms and depend to a large extent on their symbionts for prey utilization.

Fig. 1.

Longitudinal section through a typical carnivorous pitcher plant, Nepenthes × coccinea. (A) Pitcher hood with attractive glands, (B) smooth pitcher margin, (C) retention zone with loose wax crystals, (D) absorption zone with digestive glands, (E) digestive fluid, (F) hairy longitudinal ledge. Glands are drawn disproportionally large for better visibility.

Fig. 2.

pH values of fluids from different CPPs; plants grown in the greenhouse (authors' own observation), P < 0·01 (Kruskal–Wallis H-test). Outliers are marked as open circles and extreme values as asterisks.

Fig. 3.

Diversity of phytotelmata for different species of pitcher plants.

Fig. 4.

Diversity of the most important taxa of inquilines in CPPs.